S . Afr. J . Antarct. Res., Vol. 8., 1978 21

Smirh. V. R. Note> on the feeding of £cte1mwrrhinu.f similis Water­ Winterbottom and R.A. D) er. Cape Town, A.A. Balkema, 1971. ho:Jsc (Curculionidae) adults on Muion Island. Oecalof!ia, 29, Va n Zindercn Bakker, Sr.. E. M. The glacia ti on(~) of Marion Island 269-27 3. 1977b. (sub-Anta rctic). Palaeo:-cology of Aj;-ica, !!, 16 1-178. 1973. S111ith. V. R. A qualitative de,criptio:~ of energy ll.>w and nutrient Van Zind~ren B:t h.J..cr Sr.. LM. GcJ::;Jiogy of the Marion anJ cycling in the J\larion Island terre,trial eco.;)· ~lcm. Polar R!'rord. Prince b.lward !,lands (~uh-Antarctic). In Gt'll!'cological Relation.\ 18, 361-370. 1977c. between the Southern TelllfJt'rate Zonr and tlte Tropiml t\1mu/loiu.v, Smith, V. R. Animal-p l ant-~oi l nutrient relarion~hip.; on Ma.-ion edited by W. Golte. Main7, 1978. Island (sub-Antarctic). Oecalogia, 32, 239-258, 1978. Vcrwoerd. \V.J. Geolog). In \laricm and Prilll'l' £1bmrtf l~lmuls , Troll, C. Der jahre:\7eir liche Ablauf de~ Naturge,chehen~ in den edited by E. M. \311 Zindcrcn Bak"cr. J . l'vl. Winterbonom and ve rschiedenen K limaglirteln der Erde. ln Okolo~tischl' Land­ R.A. Oyer. Cape Town, A.A. Ba lJ..cnn, 1971. sdw}i~>forsch un!! und rergleichende Hodt_r:ebirgsfondumg. cdirej Vcrwoerd. W.J. & Langenegger. 0. Volc:tnological map of Marion by C. Troll. Wie;baden. Fran7 Sreincr Verlag. 1966. lsl:lnd. In Marion ami Prince Edu·m·.f lslalldS, edited by E.M. Va n Zindercn Bakker. Sr.. E. M. Quarernary climate., and An tar::ric van Zinderen Aakker, J.M. Winterbot tom and R. A. Dyer. Cape biogeography. In Amarc1ic Ecology. Vol. I. edited by M.W. To,,n, A.A. Balkema. 197 1. Holdgarc. London. Academic Pres~. 1970. William'>. D.F. & Kean). J. Compari,on of radiolarian plantonic Van Zinderen Bakkcr Sr., E. M . Introduction. l r1 Marion and Prince foraminiferal paleoeeanography of the sub-Antarctic Indian Edward lsland.1, edited by E.M. van Zindercn Bakker, J. M . Ocean. Quat. Res., 9, 71-86, 1978.

Plant ecology of Marion Island: a review V.R. S mith Institute for Environmental Sciences, University of the Orange Free State, Bloemfontein 930L.

Introduction Geographically, geologically a nd c limatologically, Marion the results and observations gai~ned from botanical and plant Is land is truly oceanic (Van Zinderen Bakker. 1978a). Situated ecologic:ll investigations carried out by various workers since in the sub-Antarctic region, it is subject to cold tempera tures 1965. The extensive limnological and phycological data are (annual mean a ir temperature 5.1 "'C). a high rainfall p resented e lsewhere in this volume and arc therefore 11ol ( > 2 500 mm per annum) and a high incidence of galc-fo.-ce considered in tl1is account. winds. T he moderating influence of the ocean . however, prevents the bitterly cold weather which characteriLes con­ The vegetation tinental winters of most northern hemisphere subpo lar sites, and permafrost does not occur. Description of the plant communities The is la nd consis ts of two distinct lava types, a grey 'Because of the rigorou:, terrestrial environment, the is land's prc-glacial and a black post-glacial e ruption. As expected of geographical isolation and its geologically recen t o rigin. only a young \ Oicanic island, the morphology of any particular 38 'ascular species occur in the island flora. of which only si\ a rea is strongly dependent upon the geological structure and contribute significantly to the aerial cover and s tanding crop most of the island surface is o f a primary construction with o f the vegetation (Smith. 1977a). An overall. oAshore view of no subsequent modification of landfonns through fluvial the isla,ld presents a bleak. barren appearance owing to the erosion. There is. therefore. a striking contrast between the absence of trees or tall shrubs in t he vegetation. However. 80 formerly g laciated a reas and those whic h have s ubsequent ly s pecies o f mosses (Van Zanten. 197 1) a nd 36 s pecies of liver­ been covered by younger. black lava fl ow~ . These latter wort~ (Grolle. 1971) occur and these form an important generally form a hummocky. ''ell-,egetated mosaic of slope component of the vegetation of ~ome a reas. hcrbfields. mi re. bog and f]aeldmark. while the s mooth Huntley (1967. 1968. 197 1) groups the island's pla nt topography o f the glaciated areas otrcrs little protection from communities into five complexes: s lope. S\\amp. salt-spray. wind ero~ion and ~uppo rts mainly !jaeldmar/.. on the ridges biotic and wind-desert. lie recognises 13 ·noda · in these and mire vegetation in the ill-drained basins. complexe:.. depending upon t he dominant s pecies present. The coastal plain on the northern. eastern and south­ The paucity of the vascular flora and the wide ecological eastern portions of the island forms an area 4-5 km wide, amplitude of many of the ~pccies prevent the classillcation of ris ing gently fro m sea-level to the foot o f the mo untai no u s the noda into narrower ca tegoric~ . However, these groups interior at about 300 m a.s.l. In contrast, the western and provide a clear general picture o f the physiognomy and ~ou thern coastal areas con~ist of a narro''· discontinuous ecology of the -.eg.ct:ttion and they arc described in detail, in­ plain of less than 100 m a lt itude and occu pied largely by cluding their noristics, distribution. edaphie, and micro­ halophytic p lant communities capable of withs tanding the climatic conditions. large amounts of sea-spray deposited onto the :.urface by the N.J .lVI. G re111mcn {unpublished) conducted a detailed strong. predo minantly wes terly wind ~. phytosociological survey of the island during 1973-75 and T he his to ry of research o n the island is well documented approximately 600 releve:l were made. A t each re leve a in Van Z inderen Bakker ( 197 1, 1978a). This paper reviews number of habitat factor:. were recorded, including a ltitude, 22 S. Afr. T. Antarkt. Nav .. Deel 8. 1978 slope. aspect, soiI type, ground water-level. drainage, soil (Huntley. 1971), as it comprises many stands intermediate water content and chemical composition. exposure to wind, between Acaena- and 8/ec/mum-dominated noda. It is not an infl uence of animals and of sea-spray. Preliminary analysis ecotonal community and often occurs independently of other of the flo ristic clata (N .J.M. G remmen, unpublished 1973/74 herbfields. Many of the communities recognised by Smith expedition report) yielded 42 plant communities which can (1976a, 1977a) as scrub or open-fernbrake are localised be arranged into the same 5 complexes mentioned by Huntley vari ants of Azorelfa herbtlelds sensu Huntley (1971). On the ( 1971). west coast of the island. however, especially near Kaalkop, (i) The slope complex luxuriant herbfield communi ties are found, consisting of On well-drained slopes, away from the immediate effects large Azorelfa cush ions, Acaena, 8/eclmum and P. cookii and of sea-spray deposition and the influence of animals, mixed these are quite distinct from open-fernbrake. herbfield communities occur, comprising ferns, cushion Azorefla-Poa montane herbfield: This nodum occurs in plants, dwarf-shrubs and grasses. Superficially and physiog­ montane sites, usually above 200 m and is best developed on nomically these communities resemble the dwarf heaths of cir1der beds on the north-western island plateau. On the ex­ the Scottish Highlands (Pearsall , 1950), the ·wet heaths' or posed crests of slopes at lower altitudes, a narrow band Gough and other cool temperate islands (Wace, 1961 , 1965) (3-5 m) of P. cookii understoried by large cushions of and the subglaeial herbfield of Macquarie Island (Taylor, A. se/ago often occurs. Both these species are tolerant of 1955). T he dominant edaphic condition governing the distri­ cold air and soil conditions and Smith ( 1976a, 1977a) regards bution of herbfield noda on the island is good drainage. these 'slope-crest" communities as low-altitude examples of Climatic factors. especially wind and temperature, are im­ Azorelft1-Poa montane herbfield. portant in determining v.hich nodum will occur at a particular (ii) The swamp complex site. The water-table is at, o r just below, the surface at all sites Bfec/uum1 penna-marina fernbrake: Variants of this group within the swamp complex. According to the nature and dominate the slope v~getation of low altitude black lava source o f the water supply, Huntley (1971) classifies the noda areas. Short, hardy 8/echnum fronds form a dense carpet, in this complex as bogs (dependent almost entirely on rain­ predominantly on slopes with a northern or eastern aspect water), mire (dependent on surface water drainage from sur­ Acaena magelfanica ( A. adscendens), Azorella se/ago, Poa rounding slopes), spring (entirely dependent on ground w.lter) cookii and Agrostis magellanica commonly occu r in th.is and flush (receiving both ground and rainwater suppl:es). nodum and Huntley ( 1971) describes how 8/eclmum is not Bogs: typical oligotrophic, ombrogenous bogs are not only limited by habitat factors within slope areas but also by common and do not form raised bogs as in the northern competition with these other species. In areas offering less hemisphere. The hepatic 8/epharidophyffwn densifolium and protection from wind, or wh ich are occupied by a large other bryophytes form a continuous carpet over the water­ number of rocky outcrops breaking the soil cover, there is logged peat and vascular vegetation cover is seldom > 10 per an increased importance of Azorefla and Acaena. Basal cent, usually comprising scattered individuals of Agrostis vegetation cover of these areas is lower than in better­ magel/anica. Momia jiuumw and Ranunculus biternatus. developed fernbrake areas and the vegetation has an open, Mires: At sites with sufficient drainage to prevent bog scrubby appearance. formation, mire occurs. dominated by A. mageffanica which In primary production and standing crop studies of the often forms a dense sward on gently sloping, lowland areas. slope plant communities, two variants of fernbrake were JwiCII.\" scheuchzerioides is common and Uncinia dikei occurs distinguished; closed fernbrake with Bfechm1111 forming a at drier sites. T he dominant bryophytes are Jamesoniella continuous (> 70 per cent) aerial cover and scrub (Smith. colorata, Drepanocladus uncinarus, Ptychomnion ringianum, 1976a) or open fern brake (Smith 1977a), where the 8/eclmum Racomirrium lanuginosum and 8/epharidophylfum densi­ cover is < 70 per cent. Closed and open fern brakes occupy folium. Phytosociologically (N.J.M. Gremmen, unpublished), approximately 18 and 22 per cent respectively of the surface a much more complicated variation in mire vegetation than area of low altitude ( < 200 m) black lava flows on the island's that described by Huntley ( 197 1) is apparent, largely because eastern coastal p lain but, together, only about I per cent o f the phytosociological technique concentrated on cover­ grey lava areas (Table 1). abundance and sociabi li ty indices and considered the bryo­ Acaena magellanica hcrbf1eld: Wet depressions and drain­ phyte component in great detail. age lines in slope areas are generally occupied by a dense Mi re and bog vegetation, as described by Huntley (1971). canopy of Acaena interspersed with long, etiolated 8/ec/mum occupy 37 and 59 per cent of the black and grey lava flows fronds and understoried by a luxuriant growth of the moss 8rachyrhecium mtabufum. This community. classified as mire Table 1 or swamp in the phytosociologi.cal study, is very similar to Percentage surface area occupied by the various plant communities the A caena mageflanica- Torrula robust a communities of on black and grey lava flows below JOO m a.s.l. on the island's sheltered depressions attracting run-off water on South eastern coastal plain. Georgia (Walton, 1973, 1976). Azorelfa se/ago herb field: A. selal{o is the most ubiquitous Community Black lava Grey lava species on the island, occurring at wet and dry sites fro1111 ( %) ( %) sea-level to the extreme altitudinal limit of vascular plant growth ( Huntley, 1970, 1972a). ft was found in 7.1 per cent Closed fernbrake 18,0 0,04 of the 457 quad rats ( I0 X 10 m) examined in 1965/66 (Huntley, Open fernbrake 21.6 1.1 197 1) and contributes up to 59 per cent of the total aerial Drainage li ne 0,3 0,0 Tussock grassland 0,6 0.4 standing crop of slope plant communities. However, recogn1- F;aeldmark 20.9 39.3 tion of A. se/ago herbfield as a nodum is based more on an Mire and bog 37.4 59.0 examination of the quadrat data than on fie ld observations S. Afr. J. Antarct. Res., Vol. 8.• 1978 23 respectively on the island's eastern coastal plain (Table I). (v) Tlte wind-desert complex Springs: the influence of ground-water on springs is Exposure to wind limits the growth of most plant species localised ar1d manifested by a distinctive community, seldom on the island's rocky plateaux. The o pen plant communities more than a few hundred square metres in area and domin­ of rocky. windswept Arctic and alpine tundra areas have lx.--cn ated by Acaena magellanica. Several other species. notably termed /jae/dmark (Warming, 1888) and this term. or its Pringlea tmti~corbutica. Poa cookii, Alomia {olllana, Ramm­ English equivalents (feldmark. fcllfield), has been applied to culus biternarus and the mosses Drepanoc/(u/us uncinatus and sub-Antarctic sites at Macquarie (Taylor, 1955), South RhyncltoStl'J:ium brttchypterygitllll are common and may Georgia (Wielgolaski, 1972a) and the Falkland islands attain optimum vigour in spring a reas. (Warming, 1888). Flush: flush vegetation occurs mainly below springs where Or1 Marion Island, fjaeldmark occurs at low altitude sites cold water spreads through the peat and an open moss turf on windswept ridges and forms the predominant vascular dominated by Bryum lael'igatum and Breutelia imegd(olia plant community above 300 m. extending up to approximately arises. The nodum is restric ted in area and dis tribution and 750 m a.s.l. A:orel/a se/ago cushio ns dorninate these areas, is usually an ecotone between spring and mire. In upland which occupy approximately 21 and 39 per cent of black and areas where ground waters a rc concentrated in steep-sloped grey lavir flows respectively below 300 m altitude on the valleys, flush communities also occur. eastern coastal plain (Table 1). Aerial cover values of A:orel/a (iii ) The salt-spray complex jjaeldmark communities below 150 m a.s.l. may be as high Rainwater-leaching limits the influence of the large amounts as 55 per cent, owing to the occurrence of epiphytic Agrostis of salt-spray deposited o nto the island by wind to a narrow, magellanica on the A:orel/a cushions (Smith. 1976a). discontinuous belt along the east coast and a wider zone on the A bryophytc-domiflated wind-desert community, termed west coast. Huntley ( 197 J) recognises two noda within the Ditricltunt-Bartramia montane desert ( H untley, 1971), ex­ vegetation of the salt-spray complex: ( I ) Tillaett moscltata tends from approximately 300 m to 1200 m a.s.l. Dominated halophytic herbfield: dominated by the crassulaceous herb by Ditric/wm strictum and Barrramia patens, the community T. moschaw. this nodum occurs at si tes experiencing very varies in size from a few moss cushions and lichens on iso­ heavy salt-spray a r1d inundation by waves. (2) Cotttla plumosa lated pebbles on the summits of the highest peaks to an herbfield; occurs in coastal .wne areas influenced by animal extensive carpet of bryophytes covering black lava flows at manuring. The composite C. pltmtosa can wi thstand heavy lower altitudes. deposition of salt-spray but requires soils manu red by animals Altiludi nal distribution of the ''ascular pla nt species to attain maximum vitality and usually occurs with other Temperature is perhaps the most important factor contri­ coprophilic species. especially Callitridte anwrctica, Momia buting to the altitudinal limitation of the distribution of .f(mta11a and Pva cookii. Huntley ( 1971) points out that this vascular plants on the islands. Thermograph recordjngs nodum is as much part of the biotic complex as it is of the ( H untley, 1970) and radiosonde data (Schulze, 197 1) indicate salt-spray complex. winter lapse rates of 0,45 °C per 100 m on the island. The (iv) Tlte biotic complex 0 penguin rookeries. seal wa ll owing grounds, albatross nests a ltitudinal range occupied by the island plants are the two and other biotic influences (Gillham. 1961: Wace, 1960. shore-zone halophytes. Tillaea mosclwta and Colt/la plumosa. 1961: G reene, 1964: Smith & Walton, 1975). Huntlcy ( 1971) Til/aea never occurs above 100 m a.s.l.. while Cotttla is only considers that the manuring influence of birds and seals i.s found above this altitude in areas heavily influenced by of more importance than salt-spray in determining the manuring. These species depend on sa lt-spray deposition or distribution of P. cookii tussock grassland on Marion Island. biotic influences for their succes~ and these factors, not wind as stands of this community arc frequently found several or temperature, are responsible for their distribution. kilometres from the coast. where salt-spray deposition is s light but where s mall petrels and prions have established Plant phenology burrows. T he importance of these birds on the nutrient Only preliminary observati ons on the phenological develop­ status of t:1e plants and soils of these areas has been described ment of the island plants are available. Huntley ( 1970) states by Smit!1 ( 1976b). that the island's vascular species may be classified into three 24 S. Afr. T. Antarkt. Nav .. Deel 8. 1978 groups according to when they begin vegetative activity and developed layer of manganese nodules. ln poorly dra ined in itiate flowering. In 1965. A. se/ago. A caena lnaxellanica and soils on less steep slopes a gleyed horizon occurs beneath the P. cookii commenced growth soon after day length began to p linthic layer and extends to bedrock. increase (Ju ly-August), long before the rapid t' ise in mean Fjcteldnwrk soils consist o f skeletal, gravelly loams of a daily air temperature for that year (November). The more shallow na ture. An A horizon occurs beneath A:orel/a cold-sensitive species. B. pen1W-1110rina, U. tlikei and A. cushions bu t, where no plants a re growing, the soil surface is magellanica. did not show signs of new growth before Novem­ eit her bare o r covered by a layer of small pebbles. Beneath ber. The otl1er major vascular species fell between these the surrace a brown gravelly loam, becoming orange-brown extremes and initiation of flowering or sporulating was also with depth and containing much scoriae, extends to bedrock, d istinctly different among the three phenological classes. usually within 10-15 cm of th.e surface. Cushions o f A::orella tend to grow toward the wind at Importance of introduced plant species in the ~·egetation low altitude fjaeldmark sites, the leeward margin dying off and Thirteen of the 38 vascular species recorded in the island exposing a humus-rich organic black clay wh ich is colonised flora are considered a liens, having been introduced by man. by other plants. Huntley ( 1971) provides 110tes on the localities at which these A typical mire peat p rofile under Agrostis magellanica alien species occurred in 1965/66 and these were updated by consists of a light-brown surface horizon approximately 20 cm a later study in 1973/75 (Gremmen, 1975). lt appears that deep overlyi ng a yellow-orange to light-brown organic clay eight alien species were introduced by sealers during the last usually extend ing to bedrock. Ir on pans and/or o rganic pans century and .five at the meteorological station since 1948. frequen tly occur in this clay layer, in which case gleying takes Four of the latter were observed flowering and collected by place and the clay is a yellow-grey colo ur beneath the ind.ura­ Huntley in 1965/66 but had disappeared from the island by tion. 1973 and can be considered as transient aliens, presumably Soil profiles under grey lava plant communities are si milar introduced with li vestock (donkeys, sheep and hens), fodder to those under corresponding black lava communj ties but or soils. No livestock currentl y exists on the island. their horizon development is genera lly poorer. The remaining alien species introduced since 1948. Rumex Three featu res render the classificati on of the island soil s. acetosella, is considered to be a persistent al ien with a very according to international systems. difficult: their immaturity, restricted distribution, occurri ng at only two local ities and the negli gible influence of parent material on the profiles, and not invading the su rrounding vegetation. although seeds and the marked enect of slight variations in topography and wind. seedlings are produced in abundance. R. acetosella is a exposure on these profiles. Taylor ( 1955) and Huntley ( 1971) widespread invader of the su b-Antarctic, being found on five classify Macquarie and Mari on Island soils respectively island grou ps. Despite successful flowering and seedling according to the most important factors influencing their establishment. the species does not seem to be in vading the development. They regard soils under slope plant communi­ natural vegetation on any of these isla nds (Wa lton. 1975} ties as higllmoor peals and those of mire and bog areas as ancl in extreme cases may be maintai ning itself only by lowmoor peals or fen and bog peals. SoUs of these groups repeated introductions. from conti.nen tal areas o f the northern hemisphere, from A lopecurus australis anes, lichens and dwarf similar to l]aeldmark soil s of the northern hemisphe re. Noting shrubs of the USSR low Arctic and s ub-Arctic (Rodin & this similarity, Huntley ( 1971) refers to the island l7aeldmark Bat.ilevich, 1967). The cryptogamic biomass in the island soils as rawmark. The use of the terms 'tundra soils· and veget:ltion b lower than that in most northern hemisphere 'dry tundra' soils in referring to s ub-Antartic jjac/dmark tundra sites. However, the large amo unt of dead material soils (Taylor, 1955) is inappropriate as they imply strongly which accumulates in the island vegetation causes the above­ gleyed mineral soils overlying permafrost. ground s tanding crop ( 12 57 1 kglha) to be substantially higher Omithogenic soil.,: True ornithogenic soils (soils essent.ially than in these northern sites. In contrast to the high above­ devoid of organic matter other than bird droppings) are ground standing crop the island vegetation possesses a lower restricted to penguin rookeries. Many o f the coastal zone q uantity (19 858 kg/ha) of below-ground plant matter than soils, however. show evidence o f having been influenced by do comparable sub-Arctic and low Arctic wndra vegctations birds (Smith, 1976c, 1977b), even if only indirectly. e.g. (23 070-62 260 kg; ha : Rodin & Batilevich, 1967). T his agrees increased rates of organic matter accumulation in the form with similar observations elsewhere in the sub-Antarctic of roots, s tems and leaves of plants whose growth is stro ngly (Jenkin & Ashton. 1970: Walton N al. 1975) and suggests stimulated by manu ring. Lindeboom ( 1979) presents evidence that the limitations to plant growth in the hars her environment that the rate of peat accumulation is strongly stimulated by of the sub-Arctic and low Arctic are relatively more the establishment of a penguin rookery in the vicinit y. The se.,erc for the above than the below-ground pans o f plants. high o rganic content, dark colour and low pH of soil s of the coastal zone at Duiker"s Point (Smith, 1978a) and the occur­ Mineral cycling rence of squid beaks in some soil profiles of this area indicate o mithogenic influences during profile development. Soil chemistry Physical and chemical weathe ring of parent material seems S tanding crop and primary production to play a minor role as a mechanism supplying nutrients to Standing crop va lues of the various plant communities the soil (Smith. 1977c). Atmospheric industrial pollution is may be found in Huntley ( 1972b) and Smith ( 1976a, 1977a). negligible a nd the major fac tors influencing plant and soil while estimates of the primary production of some of these nutrient status on the island arc the leaching eiTects of the communities arc provided in Smith ( 1978b). Summaries of high rainfall and chemical enrichment from the surrounding the standing crops of slope, mire and l]aeldmark com­ ocean, either in the form of ~alt - s pray o r as manuring by munities on the two Ja va types and of the vegetation o f the sea-going birds and mammals. This agrees with o bservati ons eastern coastal plain as a whole a rc presented in Table 2. A ll from Signy Island (Alien et al. 1967: Collins e 1 al. 1975), values a re those at approximately the time of maximum Elephant Island (Allison & Smith. 1973). South Georgia above-ground biomass. Slo pe plant communities support far (Walton & Smith. 1978) and Macquarie Island (Jcnkin. higher to tal standing crops (2 937-6 645 g; m:) than do those of 1975). Quantitative a s~ess me nt s of the mineral and energy mire (2 264-2 663 g 1m") and l]cwldmark (I 174-2 742 g m ") contributi on!> by birds to the Marion Island terrestrial eco­ areas. Plant communities on grey lava exhibit markedly lower system arc provided in several papers in this volume. standing crop va lue ~ than do those on black lava. This is H untley { 1971) found the predominant ionic order in more likely due to the higher altitude of grey lava areas a nd Mari on Island soils to be Mg> Ca > K > Na > Cl but that thei r greater expos ure to wind caused by the nat. glaciated this sequence depended on the proximity of the so il to the topography than to the difTcrences in soil fertility between shore and Na and Cl greatly predominaLe in ~oi l s of the the two lava types noted in the next section. coastal region. Grobbelaar ( 1974) found that the io nic o rde r Highest standing crops are found in slope plant communi­ in the soil solution of an area approximately 500 m inland is ties in which P. cookii and A. se/ago predominate. since these Cl > Na > Mg> Ca> K :::: Fe> SO,> Mn> Zn and corres­ s pecies have substantial accumulations o f dead matter in po nd~ clo~cly with that in the sea water ~ urrounding the their tussocks and c ushions res pectively. Jnland tussock b land. grasslands support the hi ghest above-ground biomass A summary of the !>Oil chemistry of Lhe island is pre­ (805-1 008 gj m' ) components owing to the presence of la•-ge sented in Table 3. Oetails of the analytical methods arc pro­ stems of A. magellanica. luxuriant tussocks of P. cvokii a nd vided in Smith ( 1977b). Most island soil s arc highly organic a well-developed bryophyte stratum. and the organic fracti on provides the greater part of the cati on F]aeldmar/, supports the lowest above-ground biomass adsorbing powers (Smith, 1978a). The hi gh o rganic content ( 111 -238 g/m' ) of a ll tr.e i~land communiti es (Table 2). Fve n al.;o <:ausc~ the soils to contain appreciable quantities of lower values occur at high altitude areas where truej)acldmark organic N and P. High soil acidity a nd low temperatures 26 S. Afr. T. Antarkt. Nav. . Deel 8. 1978

Table 2 Summary of standing crops in the plant communities (g m- t)

Abo,e-ground Below- Total ground standing B i omas~• Dead** Total standing crop O.M. S.C. crop Va;c. Cryp. Total

Slo pe complex Black lava 438 80-1 224- 23J 438- 1008 528- 3654 1479- -4458 2001 - 3988 4480-6645 Grey lava 310 735 70 310- 805 372- 187 1 1182- 2676 1238- 3.156 2937- 5363 Mire Black lava 11 7 219 236 303 639 2024 2663 Grey lava 97 I·H 244 242 486 1178 2264 Fjaeldmark Black lava 238 trace 238 154 1 1779 963 2742 Grey la' a Ill trace Ill 645 756 418 1174 -- Total vegetation (kg ha 1) 2528 848 3376 9196 12 57 1 19 858 32 428

•Alive; ••excluding decomposed and humidified peat constilllents.

Table 3

Chemical status of ~u r face horizons of black and grey lava soils

• • Org. mg 100 g mg 100 g

pH • o water ea Mg Na K Total C ~.:; To tal N •. Tonl P N H .1' N NO,- N

Black lava Slopes 4,0-4,7 674 315 19 7 28 13 4 I 4 3 55 21 34 15 2,00 0.60 0.65 ~ 0.26 trace- 2.9 0 - trace Fjmddmark 5,2 217 168 3 2 9 3 3 2 2 3 17 t 6 6 16 0.73 0,42 0.0 - 0.5 0,0- trace Mire 4.2- 4,3 1667 199 5 2 13 I 5 I 3 I 26 l 2 48 5 2.34 j 0.42 0.60 - 0. ~ 9 2.4 - 6,5 0,0-0,7 Grc) la va Slopes 4.3 4.8 36 1 160 11 +3 13 3 5 I I I 29 5 20 -r- 6.3 1.70 0,92 0.66 0. 15 trace- 0,7 0,0- trace /•]aeldmark 4.9 5.0 219 t 63 3 + I 5 I 4 I 2 I 14 J 2 9 t 2 0,68 1- 0,09 0,55 0,0~ 0,0 0.0 Mire 4,2-4,4 725 - 163 6 - I 5 I 5 I I I 17 3 35 5 2,08 0.27 0.60 0,17 trace 0,0- trace

n ·tard organic matter decomposition and mineralintion saturation and total N and P contents th an g rey lava soils so that soil levels of avajlable N (NH , - Nand NO,-- N) and there is evidence that the black lava parent material a re low. This is aggrava ted by inte nsive leaching. Some innuences the c hemical composition of the overlying soil to a mire peats, no tably those occurring under gelatinous mats of greater extent than in the case of the grey lava (Smith. 1978a). Nostoc commune. a cyanophyte demon~trated to fix N on the Within lava types . slo pe soils tend to contain higher amounts is la nd (Croome, 1973). a re enricl1ed in N H 4 ' N. T he poor o f exchangeable cations than do m ire peats and rawma rk. N status of the island soils is illustrated by the fact that even Rawmark possess the lowest concentrations of plant nutrients the enhanced levels of N H , N under /\'. COIIImtme (up to on both black and grey lava nows. Where plants a re growing 6,5 mgfl 00 g soil) arc m uch lower than in a s urface peat or a t t he immedia te surface the a m o unt of o rganic ma tte r in the s im il ar bulk density from a wet tundra meadow at Barro w. top few centimetres may be high, owing to the slow break­ Alaska ( 14, 1 mg' IOO g ~o il : Flint & Ge r~per . 1974). d own of plant remains due to the low temperatures prevalent R ecent, unpublished results ind icate that most of the in /Jaeldmark a reas. Fjaeldmark soil s a re, h owever, much less is land soils conta in low contents of p lant available P but a organic than a re those of mire and slo pe complexes and a re phosphorus deficiency in the soil s has not been demonstrated. al~o poor in base content. possibly due to rapid percolation T he soils possess appreciable quantities of Mg and Na of incoming ra infall through them as a res ult of their low due to salt-spray deposit io n. The excha ngeable Ca a nd Mg water-re tentio n ca pacity. concentratio ns are approximately equal o n a '~eight basis, but on an equivalent bas is Mg predominates and it appears that P la nt chemica l compositio:. the isla nd soils a re somewha t dcl1cient in Ca regard ing plant Lea f concentra tio ns o f mineral e le ments in t he isla nd gro wth and decomposition (Smith. 1976a.c). An inverse dicotyledonous. mo nocotyledo nous and bryophyte species arc relationship exists between the amo unts of exchangeable presented in Table 4. T he values are those from living material catio ns in the soil s a nd d ista nce from the sho re and coa sta[ a t a pproxima tely the time o f m aximum a bove-gro und bio mass soils contain especially high concentrations o f Mg and Na at non-manured sites. D uring the season. leaf concentrations (Smith. 1978a). Ornithogenic coastal soils are also more of Na, K. N and P decrease in most o f the s pecies. while Ca o rganic, more acid and p ossess higher cation exchange capa­ a nd M g conte nts increase in som e s pecies a nd rema in co n· cities t han soils further inland. stant in o thers. Dicotyledon values exclude Callitriche Black lava soils display greater mean percentage cation amarctica which occurs only at manured sites a nd possesses S. Afr. J. Antarct. Res ., Vol. 8., 1978 27 especially high N, P and Na contents, agreeing wi th similar throughout the season, however, and their data include o bservations from South Georgia (Smith & Walton. 1975). ln maximum values for each element. general, the dicotyledon leaves possess hj gher concentrati ons In common with plants of northern hemisphere tundra of Ca, Mg, P and N than monocotyledon leaves and bryo­ areas, K is the predominant ash element in the leaves of the phytcs contain lower concentrations of Na, K. N and P than island plants. The N content of the leaves and roots of the do the vascular species. island plants is at most only twice that of K, agreeing with the Concentrations of N and Pin the foliage of the island plants South Georgian data but in contrast to northern hemisphere are within the ranges of those reported from northern hemi­ tundra plant species. in which the concentration of N is sphere tundras (Rodin & Bazilevich, 1967: Chapin er al. normally from J to 10 times higher than that of any ash 19 75: Wielgolaski er al. 1975) and the bryophyte values are element (Rodin & Bazilevich, 1967). similar to those of bryophytes from Signy Island (Alien er al. TheCa content is lower than in more temperate herbaceous 1967). Elemental concentrations in the Marion plants arc species but substantial concentrati ons of Mg occur, probably generall y lower than in simil ar or related species on South due to the low soil Ca status and to influx of Mg from the Georgia (Walton & Smith. 1978). These authors sampled surrounding ocean. Heavy influx of Na also occurs from this

Table 4 Plam chemical composition(%).

Ca Mg Na K N p ------Dicots 0.12-0.68 0.33 0.51 0.41 - 1.80* 1,00- 1.50 2,01 - 2.91 0.13-0.25 Monocots 0,08-0, 19 0,08 0,23 0,09 0.62 1,00 1.72 0.76-2.77 0,10 0,22 Pteridophytes•• 0.43 0.71 0,28 1.55 2.36 0,24 Bryophytcs 0.19 0.83 0.17 0,43 0,06 0. 14 0, 15 O,t!7 0.47 1.55 0,03- 0.23

*Higher values of Na are for halophytic Cow/a plumosa. Other plants rarely higher than 0.8° 0 • ++ 8/edmum penna-marina only.

Table 5

1 Swnding stock of mineral elements (kg ha ) contained in the vegetation of the island's eastern coastal plain.

Above-ground ------Below- Total Biomasl. ground in ------Dead Total total ~. crop Vase. Cry p. Total O.M. above ------(i) Black lava Calcium 12,06 5, IJ 17,19 59,13 76,32 65,55 141 .87 Magnesium 12,60 4.26 16.86 43.75 60.61 74.92 135.53 Sodium 15,86 1.23 17.09 8,35 25,44 49,09 74,53 Potas~ium 38.40 9.90 48.30 13.83 62,13 149,38 21 1,51 Iron 0,26 0.39 0,65 5,34 5.99 4. 16 10, 15 Ph osphorus 5,70 1.72 7,42 12,08 19.50 39.18 58,68 T otal ash elements 84,88 22.63 107.51 142.48 249.99 382,28 632.27 Nitrogen 56,65 19.92 76.57 128.93 205,50 257.24 462.74 Total mineral elemcllls 141.53 42,55 18-t,OS 271,41 455.49 639,52 I 095,01 (ii) Grey /al'(l Calcium 2.66 2.79 5.44 14,32 19,76 25,46 45,23 M agnesium 2.74 2.73 5.46 10.44 15,90 30,60 46.50 Sodium 4.59 1,01 5.59 2,65 8.25 29.70 37,94 Po t as~ium 11,81 7.22 19,03 4,36 23,40 65.83 89,23 Iron 0. 16 0.24 0.40 2.59 2.99 1.69 4.68 Phosphorus 1.22 1,48 2.70 3,33 6,03 18,82 24,85 Total ash element~ 23.18 15.47 38,62 37,69 76,33 172.10 248.43 Nitrogen 16,23 17.34 33.58 39. 13 72.70 98,79 171.49 Total mineral elemen ts 39,41 32,81 72.20 71i.82 149.03 270,89 419.92 (iii) Eastern maswl plain Calcium 9.52 4.50 14,02 47.03 6 1.05 54,73 115.78 Magne~i u m 9,94 3.85 13.78 34.76 48.54 62.95 111 ,49 Sodium 12.82 1.1 7 13.99 6.8 1 20.80 43.85 64.65 Potassium 31,22 9,18 40.40 11 .27 51.67 126.82 178,49 Iron 0.23 0.35 0.58 4.60 5,18 3.49 8.67 Phosph orus 4,49 1.66 6. 15 9.72 15.86 33.68 49,55 Total ash clement> 68.22 20,70 88.91 114.19 203.10 325,53 528.63 Nitrogen 45.74 19.22 64.96 104.68 169.64 214,46 384.10 To tal mineral elements 11 3.96 39,92 153.87 2 18.87 372,75 539.99 912,74 28 S. Afr. T. An tarkt. Nav., Deel 8. 1978 source and the plants characteristically contain high con­ vegetation are located underground, whereas in typical centrations of this element. Plants occurring predominantly northern hemisphere tundras more than 80 per cent of the in mire and bog areas exhibit high le::tf Na contents (Smith. nutrient standing stock is contained in the below-ground 1976c) a nd this nny represent a n adaptation enabling these sphere. plants to s urvive in these nutrient-poor sites. Grobbelaar Influence of fauna on soil a nd plant chemis try ( 1974) has demonstrated that the mire soil solution contains more than six times as much Na than any other cation. Previous workers in southern subpolar regions have com­ Retranslocation and leaching effects cause standing dead mented on the enhanced vitality, cover and production of leaf material and leaf litter to exhibit lower Na, K. N and P plant communities associated with colonies of seals and concentratio ns than the corresponding living leaves. Con­ seabirds (Moseley, 1892; Skottsberg, 1912; Wace, 1960, centrations of the less mobile elements Ca. Mg and Fe are 1961 ; Gillham. 1961; Alien e1 al. 1967: Walton & Smith, substantially higher in dead than living material. in press). On Marion Island Huntley ( 1971) describes \egeta­ tion changes due to animal activity (trampling and manu­ S tanding stocks of mineral elements in the \'t~ge t a ti on ring} and outlines differences in responses of the various The amounts of mineral elements in the plant m1tter of the pla nt s pecies to this activity. various island communities have been determined (Smith. The influence of manure is vividly manifested by increased 1977b) and they are approxim:.ttely proportional to the com­ plant vitality and colour, a change in the relative proporti ons munity standing crop:;. Table 5 summ:trises the data for of the plant species present or, al heavily manured sites. in black and grey lava areas and also provides an estim:ttc of the occurrence of more nitrophilous or coprophilous plant the element standing stocks in the vegetation of the eastern species. Soils at rnanurcd sites exhibit enhanced inorganic N coastal plain. The va lues a re those at approximately the time and P levels (Table 6) and the plants possess higher contents of maximum above-ground biomass. of N and P. and. to a lesser extent, of K, Fe and Na than do Standing s tocks of Ca, Mg and N:l in the various standing planb found in areas not influenced by manuring (Croome, crop components of black lava v<:getation arc 3 to 4 times unpubli<;hed: Smith. 1976b. 1978c). greater, and those of K, t:e. P and N. 2 to 3 time.> greater lt i~ nO\\ established that the main source of N and P to than the corresponding va lues for grey lava vegetation. The the island's terrestrial ecosystem is via the dung, urine and total mineral element standing s tock in black lava vegetation guano of seJ-going animals (Burger, Lindeboom & Williams. is 2.5 t i m~s higher than that in grey lava v.::getltion. This 1978; Smith. 1976b, 197llc). Lindeboom ( 1979) provides cannot b:! ascribed to difTerenc.::s in nutrient concentration estimates of the rate of uric acid-N production in a large between plants growing on the two lava types and is due to penguin rookery and describes the chemical and micro­ the lower standing crop of the grey lava vegetation as well biological fates of this nitrogen. as to the importance of nutrient-poor communities (mire Md fjaeldmark) on grey lava flows (Table I). A tOtal of 913 kg ·ha of mir1eral elements accumulates in Conclusion the plant matter of the \Cget:ttion of the eastern co:tstal plain. Many workers have u-;cd the term tundra to describe the The living above-ground component amounts to 154 kg 1ha. vegetation and ecology of sub-Ant:trc ti c regions. Tundra is The contribution of N to the standing stock of mineral most usua ll y defined as areas with a mean annual nir tem­ elements in the a bove-ground biomass of the island vegetation pera ture below 0 ' C and where permafrost occurs. The is 42 per cent, slightly lower than that reported in the litera­ Jnternational Biological Programme adopted a more liberal ture (45-56 per cent) for comparable northern hemisphere classification (Wielgolaski. l972a, 1972b) and the I BP tundra tundra vegetations. Pota~~ium predominates amongo;t the biome site~ comprise a wide range of conditions from high ash elements in the above-ground plant biomass on the polar and alpine through low polar and subalpinc to rela­ island, fo llowed by approximately equal amounts of Cn, Na tively warm temperate bogs and oceanic moorlands and and Mg and much lower quantities or P and Fe. This con­ include two sub-Antarctic sites (Macquarie and South trasts with rnost other tundra and subpolar vcgetations where Georgia i'>lands) and one maritime Antarctic site (Signy the content of Ca is u~ually substantially higher than that of Island). The annual temperature regimes of these :,ites show Mg (Rodin & Bazi levich. 1967) and the standing stoch of a great variety of patterns (see diagram in French ( 1974); Na arc especially low, usually below those of Fe. Rosswall & Heal ( 1975) consider that the annual tempera­ There is a substantia l accumulation of Ca. Mg and Fe in ture cour:.e is more important to the biology or the individual the dead above-ground component of the island vegetation. ecosy~tcms than is the mean an11ual value. Owing to the relatively low below-ground :.tanding crop, only French ( 1974) dcmon~trared that the patterns of annual 59 per cent of the mineral elements inco rporated in the i-,land temperature CUI'\ CS for the \arious I nP tundra site~ :.how a

Table 6 lnlluence of anim

Organic N ( u 0 ) NH, N (mg lOO g) NO. N Cmg lOO gl - ·-- .\' S.D. range .\" S.D. range .\" S.D. range - Uninflucnccd soils (n 10) 2,6 0.72 1 . ~ .1.9 1,4 1.26 0 :u 0,2 0.2~ 0- 0,6 Influenced soils (n JJ) 3.6 2.51 1.7 12.1 121 .6 472.1 2.0 I 417.7 36.6 74.95 0- .150,0 E\duding \\allo" mud' (11 JOI 2,8 0.7.1 1,7 .l,9 19,7 2s,:n 2,0 11 7.5 11.3 21.61 0 1:17.8 S . Afr. J. Antarct. Res .. Vol. 8., 1978 29 gradient of warm oceanic-warm continental cold oceanic­ Everett, K.R. A survey of the soils in the region of the South cold continental. On the basis o f this s ubjective classification. Shetland Islands and adjacent parts of the Antarctic Peninsula. Marion Island fits in with the warm oceanic group comprising Institute of Polar Studies Report. 58. Ohio Stntc University. 1976. Flint, P,S. and Gcrspcr. P.L. Nitrogen nutrient levels in Arctic Macquarie Island. Glenamoy (west coast of Ireland. 54 V 12' N). Moor H ouse (U.K .. 54 65'N) and Hardangcrvidda (southern tundra soils. In Soil Oq:anisms and Decomposition in Tundra. edited b) A.J. Holding et al. 375-387. Fairbanks. Unh·ersity of Norway, 60°36' N). None of these sites experiences perma­ Alaska, 1974. frost. South Georgia a lso l:elongs to this group, although French, D.D. Classilication of JBP tundra biomc sites based on its climate is cooler than Marion or Macquarie. climate and soil properties. In Soil 0 1;rHmisms and Decomposition The vegetation physiognomy and soil chemical characteris­ in Tundra, edited by A.J. Hold ing er al. J-25, Fairbanks. Uni­ tics of these JBP warm oceanic tundra s ites arc similar to \Crsity of Alaska. 1974. those of Marion Is land. French ( 1974) also constructed a Gillham. M.E. Modifications of sub-Antarctic nora on Macquarie less subjective classification of the IBP sites using Principal l ~land by scabird~ and ~ea elephants. Pmc. R. Sor. Victoria. 74, Component Analyses of climate and soil parameters. Super­ 1-12. 1961. licial comparison of the extensive soil chemical and climatic Godley. E.J. The botany of southern Chile in relation to New data available for Mario n Isla nd (Smith, 1977b) with those caland and the sub-Antarctic. Proc. R. Soc., 8152, 457-475, 1960. used in the PCA o r the tundra sites (French, foe. sit.. BJown Grcene. S.W. Tl:e va,cula r ll ora of South Gcln nds. S. A/r. J. Sci .. 63, 235-241, 1967. occanic c lima tes of sub-Antarctic islands ( for example. see Huntley, B.J. A j/oristic and etological account of the reKefation of the thennoisopleth diagram in Van Zindcrcn Bakker. 1978a) /Harion and Princ·r £dll'r.rd lslcmds, South Indian Ocean. MSc distinguishes their ecosystem from Arctic and sub-Arctic thesis, University of Pretoria, 1968. continental regions with their extreme temperature ranges. Huntley. 8.J. Altitudinal distribution and phenology of Marion In this respect the islands have more in common with the Island \3£Cular plant~. Tyds. :vmmtrh't'l .. 10, 255-262, 1970. tropical alpine biomes which experience a characteristic Hunrley. B.J. Vegetation. In Afarion and Prince £d11'ard /.viands. diurnal c limate regime with a very slight seasonal variation edited by E.M. \an Zinderen Bakker. J.M. Winterbottom and in temperature (Van Zindcrcn Bakker, 197tlb). R.A. D)e.r, 98-160. Cape T of the international Pedo/of!.r. London. McGraw-Hill. 1941. tundra biome ~ite~. In Soil Organi.\111\ ami Drmmposition in Konono\a. 'vl.M. Soil Orf!anic ftfauer. /11 Nmure, Its Role in limtlra. edited b} A.J. Holding. 0. W. Heal. S. r. Maclean and Soil Formation all(/ in Soil Fertility (second Fngli~h translation P. W. Flanagan, 27-48. Fairbanks. Uni\ en.ity of Ala~ka. 1974. h) T.Z. NowakO\\~J..i and A.C.D. Ne\\ man). Oxford, Pergamon, Burger, A.E., Lindeboom, H.F. and William~. A.J. Mineral and 1966. energy contributions of gun no uf selected s pecie~ of birds to the Lindcboom, H.J. Clwmiml mu/ microbiolo1:iml aspects c~f the Markll1 Island ecosystem. S. A/r. .!. Anwrct. Res., 8, 5H-69, nirrog<'n c.t·cle on Maricm Island (sub-Anturffic). PhD thesis, 1978. Rijksuniversitcit of Gri.iningert. 1979. Chapin. F.S., Van Clc\e. M. and Tieszcn. L.L. Seasonal nutrient M Press. 1892. ;llpine Research. 7. 209-226, 1975. Paradakis. J. Soils of rhe World. Am~terdam, I he\ ier, 1969. Collin~. N.J., Baker. J.H. and Tilbrook. P.J. Sign) ! ~land. maritime l'ear~all. W.l-1. Mountain., and Moorlands. Lt1ndon, Collins. 1950. Antarctic. In Stmc·turr ami Function of Tundm Ecosystems. I'C

duction. In Struc/1/re and Function ofTwrdra Ecosystems, editej Winterbottom and R.A. Oyer, 1-15, Cap~ T own, A.A . Balkema. by T. Rosswall and O.W. Heal, £col. Buff. (Stockholm), 20, 197 1. 7-16, 1975. Van Zinderen Bakker, E. M. Sr. Origin a nd general ecology of the Schul ze, B.R. The climate of Marion .I sland. In Mm·ion and Prince Marion Island ecosystem S. Afr. J. Antarct. Res., 8, 12-20, £{/ward Islands, edited by E.M. van Z inderen Ba kker, .I .M. 1978a. Winterbottom and R.A. Oyer, 16-31. Cape Town, A.A. Balkema, Van Zinderen Bakker. E. M. Sr. Geoecology of the Marion and 197 1. Prince Edward Islands (s ub-Antarctic). In Erdwissenschajiliche Skottsberg, C. The vegetation in Satuh Georgia. Wiss. Ergelm. Forsclumg. 13. edited by W. Golte, Mainz, 497-515, 1978b. Schwed. Sudpo/ar - Exp. 1901 -1903. 4, (12), 19 12. Vilensky. D.G. Concerning the principles of genetic soil classifica­ Skottsberg, C. The Falkland Islands. Chronica Bot. , 7, 23-26, 1942. tion. Contriburions to the S111dy of 1he Soils of Ukrania, 6, 129-151 , Smith, V. R. Standing crop a nd nUirient status of Marion Island 1927. (sub-Antarctic) vegetation J. S. A[r. B(n. , 42, 231-263, 1976a. Wace, N.M. The botany of southern oceanic islands. Proc. R. Soc .. Smith, V.R. The efrect of burrowing species of Procellariidae on the B152, 475-490, 1960. nutrient status of inland tussock grasslands on Marion Island. Wace, N. M. The vegetation of G ough Island. £col. Monogr .. 31, J. S. A[r. Bot., 42, 265-272, 1976b. 337-367, 1961. Smith, V.R. The Nutrielll Statuses of Mm·ionlsland Plants and Soils. Wace. N.M. Vascular plants. In Biogeography and Ecology in MSc thesis, University of the O.F.S., Bloemfontein, 1976c. Anlarcrica, edited by J. va n Mieghem and P. van Oze. 201-266. Smith, V.R . Vegetation standing crop of the grey lava flows and The Hague, W. Junk, 1965. of the eastern coastal plain on Marion Island. J.S.A[r.Bol., 43. Walto n, D.W.H. Changes in standing crop and dry matter pro­ 105-1 14, 1977a. duction in an Acaena community o n South Georgia. In Primary Smith, V.R. T he chemical composition of Marion Island soils, Production and Produc1ion Processes, Tundm Biome. Proc. pla nts a nd vegeta ti on. S. A[r. J. Anlar('(. Res., 7, 28-39, 1977b. Conference IBP Tundra Biome, Dublin, April 1973, edited by Smith, Y.R. A qualitative description of energy flow and nutrient L.C. Bliss and F.E. Wielgolaski , 185-190. EdmontOn, University cycling in the Marion Island terrestrial ecosys tem. Polar Reco•·d. of Alberta Press, 1973. 18, 361-370, 1977c. Walton, D.W.H. European weeds and alien species in the sub­ Smith, V.R. Soil chemistry of Marion Island (sub-Antarctic). S. Afr. Antarctic. Weed Research, 15, 271-282, 1975. J. Sci., 74, 174-175, 1978a. Wal!on, D.W.H. Dry matter production in Acaena (Rosaceae) on a Smith, V.R. Standing crop and production estimates of selected sub-Antarctic Island. J. £col., 64, 399-4 15, 1976. Marion Island plant communities. S. Afr. J. Antarc1. Res., 8. Walton, D.W.H., Greene, D.M. a nd Callaghan, T.V. An assess­ 102- 104. 1978b. ment of primary production in a sub-Antarctic grassland on Smith, V.R. Animal- plant- soil nutrient relationships on Mario n South Georgia . Brit. Anlarct. Surv. Bu{{. , 41 & 42, 151-160, Island (sub-Antarctic). Oecologia, 32. 239-253, 197llc. 1975. Smith, R.I.L. and Walton, O.W.H. Sout h Georgia, sub-Antarctic. Wallon, D.W.H. and Smith. R.I.L. The chemical composition of In S rmcll!re and Funclion of Tundra Ecosywems, edited by T . South Georgian vegetation. Brit. An1arc1. Surv. Buff. (in press). Rosswall and O.W. Heal. £col. Bu{{. (Srocklwlm), 20, 399-423, Warming, E. On Grasslands Vegetation. Engler's Jahrb., 10, 1888. 1975. Wielgolaski, F.E. Vegetation types and primary production in Taylor, B.W. The flora, vegetation and soils of Macquarie Island. tundra. In .Proc. 4th Jut. Meeling Bioi. Prod. Tundra, edited by Aus/ralian National Awarclic Research Expedition Reports, B, F.E. Wielgolaski and T. Rosswall, 9-34, Stockholm, Berlinska 2, 1-192, 1955. Bot ryckeriet, 1972a. Thornthwaite, C.W. The climates of North America. Geog. Rev., Wielgolaski, F.E. Vegetation types and plant bi:)mass in tundra. 21. 633-654, 19 3 1. Arcric and Alpine Research. 4, 291-305, 1972b. Van Zanren, B.O. Musci. In: M(ll·ion and Prince Edward Islands, Wielgolaski. F.E., Kjelvik, S. and Kallio, P. Mineral content of edited by E. M. van Zinderen Bakkcr, J.M . Wi nrcrbc tt om a nd tundra a nd forest tundra plants in Fennoscandia. In Fenno­ R.A. Oyer, 173-227. Cape Town, A.A. .Balkema, 197 1. scam/ian Tundra Ecosyslems, Pr. I, Plams and Micro-org:mism.f, Van Zin deren Bakker, E. M. Sr. .Introduction. In Marion and Prince edited by F.E. Wiclgolaski , 316-332, Heidelberg. Springer­ Edward Islands, edited by E.M. van Zinderen Bakkcr, J.M. Vcrlag, 1975.

Ornithological research at the Prince Edward islands: a review of progress W .R. Siegfried Percy FitzPatrick fnstitute of African Ornithology, University of Cape Town, Rondebosch 7700,

Captain Cook, who sighted but did not land at the Prince 'ollicially" discovered by the French explorer, Marion Edward islands in 1776, mentions seeing 'penguins and Dufresne, in 1772. However, it is likely th~t D ufresne was shags, the former so numerous that the rocks seemed covered preceded by skipper Ham, of the Dutch galleon Maerseveen , with them a s with a crust'. This appears to be the earliest in 1663 ( Leupe, 1868). h any event, it was H:t rris, a ship's refere11ce to the birds on these islands which had been engineer and sealer, who made the first detailed observatior1s